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98 CHAPTER 5 Ecosystems: Concepts and Fundamentals<br />

The bog also fills in from the bottom <strong>as</strong> streams<br />

carry fine particles of clay into it (Figure 5.17b, c). At<br />

the shore, the floating mat and the bottom sediments<br />

meet, forming a solid surface. But farther out, a “quaking<br />

bog” occurs. You can walk on this mat; and if you<br />

jump up and down, all the plants around you bounce<br />

and shake, because the mat is really floating. Eventually,<br />

<strong>as</strong> the bog fills in from the top and the bottom,<br />

trees grow that can withstand wetter conditions—such<br />

<strong>as</strong> northern cedar, black spruce, and balsam fir. The formerly<br />

open-water bog becomes a wetland forest. If the<br />

bog is farther south, it may eventually be dominated by<br />

beech and maple, the same species that dominate the<br />

late stages of the dunes.<br />

Old-Field Succession<br />

In the northe<strong>as</strong>tern United States, a great deal of land<br />

w<strong>as</strong> cleared and farmed in the 1<strong>8th</strong> and 19th centuries.<br />

Today, much of this land h<strong>as</strong> been abandoned for farming<br />

and allowed to grow back to forest (Figure 5.18). The first<br />

plants to enter the abandoned farmlands are small plants<br />

adapted to the harsh and highly variable conditions of a<br />

clearing—a wide range of temperatures and precipitation.<br />

As these plants become established, other, larger plants enter.<br />

Eventually, large trees grow, such <strong>as</strong> sugar maple, beech,<br />

yellow birch, and white pine, forming a dense forest.<br />

Since these three different habitats—one dry (the<br />

dunes), one wet (the bog), and one in between (the old<br />

field)—tend to develop into similar forests, early ecologists<br />

believed that this late stage w<strong>as</strong> in fact a steady-state<br />

condition. They referred to it <strong>as</strong> the “climatic climax,”<br />

meaning that it w<strong>as</strong> the final, ultimate, and permanent<br />

stage to which all land habitats would proceed if undisturbed<br />

by people. Thus, the examples of succession were<br />

among the major arguments in the early 20th century<br />

that nature did in fact achieve a constant condition, a<br />

steady state, and there actually w<strong>as</strong> a balance of nature.<br />

We know today that this is not true, a point we will<br />

return to later.<br />

Coral Reef Succession<br />

Coral reefs (Figure 5.19) are formed in shallow<br />

warm waters by corals, small marine animals that live in<br />

colonies and are members of the phylum Coelenterata,<br />

which also includes sea anemones and jellyfishes. Corals<br />

have a whorl of tentacles surrounding the mouth,<br />

and feed by catching prey, including planktonic algae,<br />

<strong>as</strong> it p<strong>as</strong>ses by. The corals settle on a solid surface and<br />

produce a hard polyp of calcium carbonate (in other<br />

words, limestone). As old individuals die, this hard<br />

material becomes the surface on which new individuals<br />

establish themselves. In addition to the coelenterates,<br />

other limestone-shell-forming organisms—algae,<br />

corals, snails, urchins—live and die on the reef and are<br />

glued together primarily by a kind of algae. 16 Eventually<br />

a large and complex structure results involving many<br />

other species, including autotrophs and heterotrophs,<br />

creating one of the most species-diverse of all kinds of<br />

ecosystems. Highly valued for this diversity, for production<br />

of many edible fish, for the coral itself (used in<br />

various handicrafts and arts), and for recreation, coral<br />

reefs attract lots of attention.<br />

Succession, in Sum<br />

Even though the environments are very different,<br />

these four examples of ecological succession—dune, bog,<br />

old field, and coral reef—have common elements found<br />

in most ecosystems:<br />

1. An initial kind of autotroph (green plants in three<br />

of the examples discussed here; algae and photosynthetic<br />

bacteria in marine systems; algae and photosynthetic<br />

bacteria, along with some green plants in some<br />

freshwater and near-shore marine systems). These are<br />

typically small in stature and specially adapted to the<br />

unstable conditions of their environment.<br />

FIGURE 5.18 Old-growth e<strong>as</strong>tern deciduous forest.<br />

FIGURE 5.19 Hawaiian coral reef.

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